TF and cancer: As base damage (e.g. oxidative, alkylation, uracil, and AP sites) has long been recognized as an important source of DNA mutations in human cancers (Maynard et al., 2009; Tubbs and Nussenzweig, 2017), the interplay between TF binding, base damage formation, and BER revealed by our study has important implications for understanding mutations in gene regulatory regions.